Taxonomy, Transitional Forms,
and the Fossil Record
Keith B. Miller
Department of Geology
Kansas State University, Manhattan, KS 66506
The recognition and interpretation of patterns in the fossil record require an awareness of the limitations of that record. Only a very small fraction of the species that have lived during past geologic history is preserved in the rock record. Most marine species are soft-bodied, or have thin organic cuticles, and are essentially unpreservable except under the most extraordinary conditions. ...
Introduction
The fossil record provides persuasive evidence for macroevolutionary change and common descent. ...
Nature of the Fossil Record
There are two opposite errors which need to be countered about the fossil record: (1) that it is so incomplete as to be of no value in interpreting patterns and trends in the history of life, and (2) that it is so good that we should expect a relatively complete record of the details of evolutionary transitions within most lineages.
What then is the nature of the fossil record? It can be confidently stated that only a very small fraction of the species that once lived on Earth has been preserved in the rock record and subsequently discovered and described by science...
Limits of the Fossil Record
Soft-bodied or thin-shelled organisms have little or no chance of preservation, and the majority of species in living marine communities are soft-bodied. ...
Potential of Fossil Record for Understanding Evolutionary Change
Given the limitations and biases discussed above, what should be expected from the fossil record? The situation is not as bleak as it may appear from my previous comments. Exceptional deposits, such as the Burgess Shale, Solnhofen Limestone, and Green River Shale, do provide surprisingly detailed glimpses of once living communities. These rare cases of exceptional preservation (fossil lagerstätten) are essentially snapshots in the history of life and are invaluable in gaining a more comprehensive picture of ancient communities. They also provide some of the most detailed anatomical data.
...
Taxonomy and Transitional Forms
Taxonomy, the process of classifying living and fossil organisms, produces its own patterns which order the diversity of life. It is thus important to recognize that names do much more than describe nature: they also interpret it. There is considerable ferment now within the field of taxonomy because of conflicting philosophies of classification, and different perceptions of which patterns in the history of life should be reflected in the taxonomic hierarchy (Eldredge & Cracraft, 1980; Schoch, 1986). Higher taxa can be either artificial groupings of species with similar morphologies (evolutionary grades), or "natural" groups sharing derived characteristics inherited from a common ancestor (monophyletic taxa or clades).
The Linnean classification system is hierarchical, with species grouped into genera, genera into families, families into orders, etc. This system reflects the discontinuity and hierarchy observed among living organisms. However, "this system leads to the impression that species in different categories differ from one another in proportion to differences in taxonomic rank" (Carroll, 1988, p. 578). This impression is false. ...
Based on the above discussion, a transitional form is simply a fossil species that possesses a morphology intermediate between that of two others belonging to different higher taxa. Such transitional forms commonly possess a mixture of traits considered characteristic of these different higher taxa. ...
Conclusions
From this brief survey of fossil vertebrates, it is clear that transitional forms between higher taxa are common features of the fossil record. The morphology of species within a higher taxonomic group becomes less divergent toward the point of origin of that group. Morphological diversity and disparity increase with time. In addition, transitional species possess mixtures of morphologic characters from different higher taxa often to the extent that their taxonomic assignment is uncertain. This pattern is obscured by taxonomy which gives a false impression of discontinuity.
...
Bibliography
Ahlberg, P.E. and Milner, A.R., 1994, The origin and early diversification of tetrapods: Nature, vol. 368, p. 507-14.
Aldridge, R.J. and Purnell, M.A., 1996, The conodont controversies: Trends in Ecology and Evolution, vol. 11, p. 463-8.
Bakker, R.T., 1975, Dinosaur renaissance: Scientific American, vol. 232, p. 5878.
Benton, M.J., 1991, Amniote phylogeny: IN, H.-P. Schultze and L. Trueb, (eds.), Origins of the Higher Groups of Tetrapods: Controversy and Consensus, Comstock Publishing Associates, Ithaca, p. 317-30.
Berta, A., 1994, What is a whale?: Science, vol. 263, p. 180-1.
Briggs, D.E.G., Clarkson, E.N.K., and Aldridge, R.J., 1983, The conodont animal: Lethaia, vol. 16, p. 1-14.
Carroll, R.L., 1970, The earliest known reptiles: Yale Scientific Magazine, October issue, p. 16-23.
Carroll, R.L., 1988, Vertebrate Paleontology and Evolution: W.H. Freeman & Co., New York, 698 p.
Carroll, R.L., 1991, The origin of reptiles. IN, H.-P. Schultze and L. Trueb (eds.), Origins of the Higher Groups of Tetrapods: Controversy and Consensus, Comstock Publishing Associates, Ithaca, p. 331-53.
Chiappe, L.M., 1995, The first 85 million years of avian evolution: Nature, vol. 378, p. 349-55.
Crompton, A.W. and Parker, P., 1978, Evolution of the mammalian masticatory apparatus: American Scientist, vol. 66, p. 192-201.
Denton, M, 1985, Evolution: A Theory in Crisis: Adler & Adler, Bethesda, MD, 368 p.
deRicqlés, A.J., 1974, Evolution of endothermy: histological evidence: Evolutionary Theory, vol. 1, p. 51-80.
Desui, M., 1991, On the origins of mammals. IN, H.-P. Schultze and L. Trueb (eds.), Origins of the Higher Groups of Tetrapods: Controversy and Consensus, Comstock Publishing Associates, Ithaca, p. 570-97.
Dodson, P., 1985, International Archaeopteryx Conference: Journal of Vertebrate Paleontology, vol. 5, p. 177-9.
Eldredge, N. and Cracraft, J., 1980, Phylogenetic Patterns and the Evolutionary Process: Columbia University Press, New York, 349 p.
Erwin, D.H., 1993, The Great Paleozoic Crisis: Life and Death in the Permian: Columbia University Press, New York, 327 p.
Evander, R.L., 1989, Phylogeny of the family Equidae. IN, D.R. Prothero and R.M. Schoch (eds.), The Evolution of the Perissodactyls: Oxford University Press, New York, p. 109-27.
Gingerich, P.D., 1976, Paleontology and phylogeny: Patterns of evolution at the species level in early Tertiary mammals: American Journal of Science, vol. 276, p. 1-28.
Gingerich, P.D., Raza, S.M., Arif, M., Anwar, M., and Zhou, X., 1993, Partial skeletons of Indocetus ramani (Mammalia, Cetacea) from the lower middle Eocene Domanda Shale in the Sulaiman Range of Punjab (Pakistan): Contributions from the Museum of Paleontology, University of Michigan, vol. 28, p. 393-416.
Gingerich, P.D., Raza, S.M., Arif, M., Anwar, M., and Zhou, X., 1994, New whale from the Eocene of Pakistan and the origin of cetacean swimming: Nature, vol. 368, p. 844-7.
Hecht, M.K., Ostrom, J.H., Viohl, G., and Wellnhofer, P. (eds.), 1985, The Beginnings of Birds: Proceeding of the International Archaeopteryx Conference, Eichstatt, 1984: Bronner & Daentler, Eichstatt.
Hopson, J.A., 1991, Systematics of the nonmammalian Synapsida and implications for patterns of evolution in synapsids. IN, H.-P. Schultze and L. Trueb (eds.), Origins of the Higher Groups of Tetrapods: Controversy and Consensus, Comstock Publishing Associates, Ithaca, p. 635-93.
Hopson, J.A., 1994, Synapsid evolution and the radiation of non-eutherian mammals. IN, D.R. Prothero and R.M. Schoch (eds.), Major Features of Vertebrate Evolution, Short Courses in Paleontology, No. 7: Paleontological Society, Knoxville, p.190-219.
Hotton, N, III, 1991, The nature and diversity of synapsids: Prologue to the origin of mammals. IN, H.-P. Schultze and L. Trueb (eds.), Origins of the Higher Groups of Tetrapods: Controversy and Consensus, Comstock Publishing Associates, Ithaca, p.598-634.
Mader, B.J., 1989, The Brontotheriidae: A systematic revision and preliminary phylogeny of North American genera. IN, D.R. Prothero and R.M. Schoch (eds.), The Evolution of the Perissodactyls: Oxford University Press, New York, p. 109-27.
McFadden, B.J., 1988, Horses, the fossil record, and evolution: Evolutionary Biology, vol. 22, p. 131-58.
McFadden, B.J., 1992, Fossil Horses: Systematics, Paleobiology, and Evolution of the Family Equidae: Cambridge University Press, Cambridge, 369 p.
McNab, B.K., 1978, The evolution of endothermy in the phylogeny of mammals: American Naturalist, vol. 112, p. 1-21.
Mikulic, D.G., Briggs, D.E.G., and Kluessendorf, J., 1985, A Silurian soft-bodied fauna: Science, vol. 228, p. 715-7.
Morell, V., 1997, The origin of birds: the dinosaur debate: Audubon, vol. 99, no. 2, p. 36-45.
Norman, D., 1985, The Illustrated Encyclopedia of Dinosaurs: Crescent Books, New York, 208 p.
Norman, D., 1994, Prehistoric Life: The Rise of the Vertebrates: Macmillan, New York, 246 p.
Novas, F.E. and Puerta, P.F., 1997, New evidence concerning avian origins from the Late Cretaceous of Patagonia: Nature, vol. 387, p. 390-2.
Ostrum, J.H., 1979, Bird flight: How did it begin?: American Scientist, vol. 67, p. 46-56.
Ostrum, J.H., 1994, On the origin of birds and of avian flight. IN, D.R. Prothero and R.M. Schoch (eds.), Major Features of Vertebrate Evolution, Short Courses in Paleontology, No. 7: Paleontological Society, Knoxville, p.160-77.
Prothero, D.R., Guerin, C., and Manning E., 1989, The history of the Rhinocerotoidea. IN, D.R. Prothero and R.M. Schoch (eds.), The Evolution of the Perissodactyls: Oxford University Press, New York, p.321-40.
Prothero, D.R. and Schoch, R.M., 1989, Origin and evolution of the Perissodactyla: Summary and synthesis: IN, D.R. Prothero and R.M. Schoch (eds.), The Evolution of the Perissodactyls: Oxford University Press, New York, p.504-29.
Radinsky, L.B., 1979, The early evolution of the Perissodactyla: Evolution, vol. 23, p. 308-28.
Romer, A.S., 1966, Vertebrate Paleontology: University of Chicago Press, Chicago,
468 p.
Schoch, R.M., 1986, Phylogeny Reconstruction in Paleontology: Van Nostrand Reinhold Company, New York, 351p.
Schultze, H.-P., 1991, A comparison of controversial hypotheses on the origin of tetrapods. IN, H.-P. Schultze and L. Trueb (eds.), Origins of the Higher Groups of Tetrapods: Controversy and Consensus, Comstock Publishing Associates, Ithaca, p. 29-67.
Stanley, S.M., 1974, Relative growth of the titanothere horn: a new approach to an old problem: Evolution, vol. 28, p. 447-57.
Stanley, S.M., 1979, Macroevolution: Pattern and Process: WH. Freeman & Company, San Francisco, 332 p.
Thewissen, J.G.M., Hussain, S.T., and Arif, M., 1994, Fossil evidence for the origin of aquatic locomotion in archaeocete whales: Science, vol. 263, p. 210-2.
Valentine, J.W., 1992, The macroevolution of phyla. IN, J.H. Lipps and P.W. Signor (eds.), Origin and Early Evolution of the Metazoa: Plenum Press, New York, p. 525-53.
Vorobyeva, E. and Schultze, H.-P., 1991, Description and systematics of panderichthyid fishes with comments on their relationship to tetrapods. IN, H.-P. Schultze and L. Trueb (eds.), Origins of the Higher Groups of Tetrapods: Controversy and Consensus, Comstock Publishing Associates, Ithaca, p. 68-109.